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Curr Top Behav Neurosci. Author manuscript; available in PMC 2016 December 01.
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Curr Top Behav Neurosci. 2016 ; 27: 1–12. doi:10.1007/7854_2015_402.

The Behavioral Neuroscience of Motivation: An Overview of

Concepts, Measures, and Translational Applications
Eleanor H. Simpson and
Department of Psychiatry, New York State Psychiatric Institute, Columbia University, New York,
NY, USA

Peter D. Balsam
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Department of Psychiatry, New York State Psychiatric Institute, Psychology Departments of

Barnard College and Columbia University, New York, NY, USA
Eleanor H. Simpson: es534@cumc.columbia.edu; Peter D. Balsam: Balsam@columbia.edu

Abstract
Motivation, defined as the energizing of behavior in pursuit of a goal, is a fundamental element of
our interaction with the world and with each other. All animals share motivation to obtain their
basic needs, including food, water, sex and social interaction. Meeting these needs is a requirement
for survival, but in all cases the goals must be met in appropriate quantities and at appropriate
times. Therefore motivational drive must be modulated as a function of both internal states as well
as external environmental conditions. The regulation of motivated behaviors is achieved by the
coordinated action of molecules (peptides, hormones, neurotransmitters etc), acting within specific
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circuits that integrate multiple signals in order for complex decisions to be made. In the past few
decades, there has been a great deal of research on the biology and psychology of motivation. This
work includes the investigation of specific aspects of motived behavior using multiple levels of
analyses, which allows for the identification of the underpinning neurobiological mechanisms that
support relevant psychological processes. In this chapter we provide an overview to the volume
“The Behavioural Neuroscience of Motivation”. The volume includes succinct summaries of; The
neurobiology of components of healthy motivational drive, neural measures and correlates of
motivation in humans and other animals as well as information on disorders in which abnormal
motivation plays a major role. Deficits in motivation occur in a number of psychiatric disorders,
affecting a large population, and severe disturbance of motivation can be devastating. Therefore,
we also include a section on the development of treatments for disorders of motivation. It is hoped
that the collection of reviews in the volume will expose scientists to a breadth of ideas from
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several different subdisciplines, thereby inspiring new directions of research that may increase our
understanding of motivational regulation and bring us closer to effective treatments for disorders
of motivation.

Keywords
Motivation; Cost-benefit analysis; Addiction; Apathy; Translational research

Correspondence to: Eleanor H. Simpson, es534@cumc.columbia.edu.

 Simpson and Balsam Page 2

1 Why Motivation Is Important to Understand

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Understanding what drives motivated behavior in humans is a truly fascinating endeavor. But
as important as our curiosity for knowing what drives us as individuals, and what supports
individual differences in levels of motivation among our friends and colleagues, is the
critical question; why do motivational processes get disrupted when the clinical and personal
consequences can be so devastating? As we will see across this volume, motivated behaviors
involve biological and psychological processes that have undergone evolution at numerous
levels, from individual molecules all the way to species-specific social organization. While
motivational processes represent heritable traits of fitness, humans suffer from a number of
disorders of motivation that can be organized into two distinct categories. The first category
is composed of the apathy and pathological deficits in motivation commonly seen in patients
with schizophrenia and affective disorders. The second category involves problematic
excesses in behavior including addictions, the pathological misdirection of motivation.
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Developing treatments for disorders of motivation requires a detailed understanding of how

motivated behavior occurs, how it is dynamically regulated under normal conditions, and
how it is disrupted in disease. This volume provides reviews of recent research in each of
these areas.

2 What We Mean by the Word Motivation

The concept of motivation is a useful summary concept for how an individual’s past history
and current state interact to modulate goal-directed activity. In this book, the authors
examine the motivation to pursue many different goals. One general aspect of motivated
behaviors is that they lead to a goal and obtaining the goal is rewarding. Thus, motivation,
defined as the energizing of behavior in pursuit of a goal, is a fundamental property of all
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deliberative behaviors. One of the earliest psychological theories of motivation, Hull’s drive
theory, posited that behaviors occur to reduce biological needs, thereby optimizing the
organism’s potential for survival (Hull 1943). However in Hull’s theory, motivational drive
functioned solely to energize responding, drive was not responsible for initiating, or
maintaining the direction of action. Later, motivation was conceptualized to consist of both a
goal-directed, directional component and an arousal, activational component (Duffy 1957;
Hebb 1955). This is the framework of motivation still in use, such that if motivation were a
vector—its length would represent the amplitude, or intensity of pursuit, and the angle of the
vector would represent its focus on a specific goal. In this analogy, a motivation vector
affected by apathy might have a reduced length in all directions and a motivation vector
affected by addiction might have an increased length and a less flexible direction. The
chapters in this volume explicitly acknowledge that motivation affects which responses
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occur as well as the vigor of those responses. It appears that we are just beginning to
understand that these two aspects of motivation have both common and distinctive neural
underpinnings. For example, circadian factors may energize the general motive of seeking
food or mate (Antle and Silver, this volume), but the specific actions that occur in pursuit of
these goals are regulated by different substrates (see Caldwell and Alders, Woods and Begg,
Magarinos and Pfaff, all in this volume). Similarly, local cues that signal food availability
may energize many food seeking actions, signals for specific foods differentially energize
actions associated with obtaining the specific outcome. Again, the neural substrates of the

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general and specific effects are somewhat distinctive. At each level, whole classes of specific
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actions are made more or less likely by these factors (Neuringer and Jensen 2010). We
suggest that there is generally a hierarchical structure to motivation in the sense that general
arousal factors such as sleep–wake cycles will affect many different motives, that activation
of specific motives (e.g., hunger, thirst, social motives) can activate many specific actions
that could lead to many specific outcomes within a general class of goals, and that more
temporally and situationally specific factors determine the specific actions that occur in
pursuit of that goal (Timberlake 2001). With this in mind, it is clear that disruptions in
motivation can occur at multiple levels of control which suggests there may be multiple
interacting ways to attempt to treat disruptions.

3 A Simplified Overview of How Motivation Might Work in the Brain

Many different factors influence motivation, including the organism’s internal physiological
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states, the current environmental conditions, as well as the organism’s past history and
experiences. In order for all these factors to influence motivation, information about them
must be processed in a number of ways; it must be evaluated and encoded, and unless the
motives are novel, the valuation and encoding will be affected by learning and retrieval
processes. A simplified overview of how such diversity of information must be processed
and integrated to result in motivation (both response selection and action vigor) is shown in
Fig. 1. Here, we organize the problem into a single, highest order concept that motivated
behaviors represent the actions associated with the highest net value that results from a cost–
benefit analysis that encompasses all of the potential influencing factors and processes.

4 Cost–Benefit Computation as the Arbiter of Motivated Behavior

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The costs associated with behavioral action may include physical effort, mental effort, time,
loss of potential opportunities, discomfort, and danger (the risk of pain and potential death).
The benefits associated with behavioral action might include fulfilling physiological and
psychological needs, obtaining reinforcement secondary to those needs, escaping from harm,
or avoidance of some of the costs listed above. As mentioned above, information entering
the cost–benefit computation for any specific motive will be processed in several ways. The
value of every cost and every benefit must be calculated and encoded. The concept of
encoding value and experimental methods for measuring encoded value are discussed in
detail by Redish et al., in this volume. It is important to consider that value must be encoded
when a goal is obtained and then stored for future retrieval when obtaining that goal again
becomes relevant. When that happens in the current moment, the assessment of value must
be conditioned both on this past experience as well as the current state and environmental
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conditions. Goals are likely to be obtained with some temporal distance from the initiation,
or even conclusion of behavioral output. Single neuron activity in several brain regions
including orbitofrontal cortex, anterior cingulate, and basolateral amygdala has been shown
to correlate with reward prediction and this work is reviewed by Bissonette and Roesch in
this volume. The encoded values of costs and benefits do not belong in absolute scales
because the values of all costs and benefits are rendered relative to the animal’s current
physiological state as well as the current conditions of the surrounding environment.

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Much has been learned about the role of dopamine in reinforcement learning, and its impact
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on motivated behavior from experimental manipulations of the dopamine system in rodents.

This work is comprehensively reviewed by Salamone et al. in this volume. In addition to
learning about the costs and benefits of a particular action, subjects also learn about specific
signals that are associated with obtaining particular goals. Such signals can have an
enormous influence on motivated behavior, and several chapters in this volume provide
details of when and how environmental cues can influence response selection and response
activation. These include Corbit and Balleine’s chapter on learning and motivational
processes contributing to Pavlovian–instrumental transfer and John O’Doherty’s chapter on
the neural substrates of motivational control in humans. Cue learning is also discussed in the
context of motivational disorder, in the chapters by Meyer et al. and Barrus et al., that deal
with substance abuse disorders and gambling.

Another aspect of the computation that energizes specific action has to do with signals that a
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particular goal is currently available. These signals occur on multiple timescales. Specific
times of day can become associated with the opportunity to obtain specific goals, and
discrete cues can signal the opportunity to achieve a goal as well as what specific ways there
are to achieve it. For example, when meals occur at a regular time of day, there are
behavioral, hormonal, and neural changes that occur in anticipation of a meal time that give
rise to the motivation to seek food (Antle and Silver, in this volume). Encountering a
restaurant can activate the specific behavioral sequences that lead to the ordering of food and
the specific foods themselves can activate specific consummatory responses. All along this
sequence of temporally organized behavior, there are concomitant changes in hormonal and
neural states that energize and guide action (Woods and Begg in this volume).

After effective encoding of all the relevant costs and benefits, a computational process (the
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cost–benefit computation) is required to resolve the appropriate direction and vigor of action
to be taken. This complex interplay of factors and processes is schematized in Fig. 1.

How conceptually the cost–benefit computation is made is currently unclear. It is still

unknown whether the value of costs and benefits are calculated on the same scale or not,
whether their weights are integrated or subtracted such that, for example, the amount of
predicted effort reduces the value of the predicted reward. Or perhaps, there is a circuit
component that acts as a comparator of these two component values. An additional
complication is that for any given motive, there are often multiple types of costs and
potentially multiple types of benefits involved because many different types of control
systems and circuits are at play (e.g., neuroendocrine, circadian, Pavlovian). This leads to
the question of how so much diversity of information can all be used to make an appropriate
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response selection and determine action vigor. Do all factors enter into a singular, highly
complex equation, as our simplified diagram (Fig. 1) may seem to imply? Or do some
systems continually run in parallel, with behavioral output as the result of a hierarchical
switching from one system to another? Or perhaps there is fluctuation in the degree to which
different factors influence the computation, e.g., the relative weights of physical and mental
costs depend on the energy state of the organism.

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Furthermore, it is possible that these alternative regulatory schemes are not mutually
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exclusive. For a detailed discussion on the potential mechanisms by which multiple

deliberative processes that are running in parallel may each influence motivation (see Redish
et al. in this volume). In the case of appetitive conditioning, there is evidence to suggest that
animals can rapidly switch between responding that is driven by two different control
systems, goal directed or habitual (Gremel and Costa 2013). Audiovisual cues can trigger
the rapid switching, implying that these two alternate circuits are constantly online and
available in parallel. On the other hand, instead of a multi-tiered, hierarchical, or switching
system, other work suggests that all information enters a singular computation process, and
the output of this meta-computation is what drives motivation. This concept is favored by
Magarinos and Pfaff (this volume) whose work on the sexual motivation of female rodents
may suggest that for this specific motive, at least some factors may be integrated into a
single decision-making process.
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Above we have described the complex situation of many different factors influencing a
single motive. It must also be recognized that at any given time, there may be competition
for multiple goals and that imbalances in the strength of the motivations for each goal can
cause conflict and dysfunctional behavior. The chapter by Cornwell et al. describes how
human well-being depends not only on satisfying specific motives, but also on ensuring that
motives work together such that no individual motive is too weak or too strong. It is
becoming clear that different motivational systems have control elements that are unique to
each system but that there may also be common substrates, perhaps close to the final steps
that determine behavioral output. This is well illustrated in the chapter on defensive
motivation by Campese et al. and in the chapter on social motivation by Caldwell and
Alders. Again, the neurobiological mechanism whereby different motive systems interact is
an important but not yet well-understood problem. Of particular interest will be to
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understand how defensive motivations interact with appetitive ones. The vast majority of
modern work on motivation concerns itself with the mechanisms of appetitive motivation.
Campese et al. show how to leverage what is now known about fear learning to understand
the neurobiological mechanisms of defensive motivations. In a similar vein, Cornwell et al.
argue for the importance of understanding how promotion/prevention motives in humans is
an important modulator of other motives. Hopefully, the future will include a greater focus
on understanding defensive motivations.

5 Research Approaches to Understanding Motivation

To increase our understanding of motivation in the brain, there are numerous approaches that
can be taken. In this volume, many different academic approaches are represented as the
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research reviewed includes clinical, experimental, and comparative psychology; and several
neuroscience subfields including, cognitive, molecular, cellular, behavioral, and systems
neuroscience. This means that specific questions or single hypotheses can be, and often are
being, approached at multiple levels of analysis. Indeed, it is when research programs
combine a number of techniques, or use information derived from a few different techniques
to propose (and test) new hypotheses that the most compelling results are obtained. For
example, the work described by O’Doherty in this volume includes the use of human fMRI
studies to investigate potential action-value signals that have been proposed from rat

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electrophysiological recordings. The research described by Redish et al. considers

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computational models of decision making and tests these models by measuring neuronal
activity during deliberative behavior. In the chapter by Ward, the approach to testing
motivational deficits in mice has very much been informed by the data from molecular and
clinical studies in humans. By phenocopying in mice the molecular changes that have been
detected in patients using PET imaging techniques, the behavioral consequences can be
probed under well-controlled conditions. In a similarly translational manner, the research
described by Robinson et al. in this volume applies electrophysiological and optogenetic
techniques in rodents to probe behaviors that are altered in people with addictions. In the
chapter by Barrus et al., the authors discuss the development of rodent paradigms designed
to test various psychological theories of substance use and gambling disorders.

6 Organismal Level Biology Is Critical to Understanding Motivation

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When multiple levels of analyses are used to investigate motivational processes, a critically
important concept becomes apparent. While the evolution of traits that support motivation
occurs at the level of molecules, proteins, cells, and circuits, it is the entire organism, and its
interaction with the environment that is selected. An example of this concept is easily seen
in the research on circadian modulation of motivation (Antle and Silver) and in the work on
motivation for eating (Woods and Begg). For example, in the case of feeding we know many
of the molecules and circuits involved in both the intrinsic, homeostatic factors which drive
the motivational to eat, such as hormones and peptides, and we also know the
neuromodulators and circuits that are responsible for some of the extrinsic/environmental
influences on eating such as predictive cues. We are beginning to understand how these
signals are integrated in order for decisions to be made and behavioral responses to occur;
though as described above, understanding the mechanism of integration is currently a critical
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area of research.

7 Motivation Gone Wrong

Patients with many different psychiatric diagnoses may experience deficits in motivation,
including depression, schizophrenia, bipolar disorder, PTSD, and anxiety disorders. In this
volume, we focus on the neurobiology of motivational deficits in depression and
schizophrenia primarily because these are the two illnesses in which pathological deficits in
motivation play a major role in patient functioning and clinical outcome (Barch et al. 2014;
Strauss et al. 2013). As such, far more research has been done on motivation in depression
and schizophrenia than any other illness. In the last few decades, it has been recognized that
the motivational deficits in Schizophrenia and depression share similarities, but also distinct
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differences. These differences occur because as mentioned above, there are many
components involved in motivated behavior and each of them represent potential
vulnerabilities that may be involved in different pathophysiological mechanisms. An
excellent review of the similarities and differences in mechanisms underlying motivational
deficits in depression and schizophrenia is provided by Barch et al. in this volume. The
central difference in these types of pathologies is that many depressed patients suffer from
impairments of in-the-moment hedonic reaction. Such anhedonia can diminish an
individual’s capacity for anticipation, learning, and effort. In contrast, patients with

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schizophrenia demonstrate relatively intact in-the-moment hedonic processing. Instead,

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patients suffer impairments in other components involved in translating reward experience to

anticipation and action selection.

There are also separate chapters that go into more specific detail for each of these
pathophysiological conditions. An update on candidate pathomechanisms for motivational
deficits in depression is provided by Treadway’s chapter. This volume devotes two chapters
to the topic of motivation deficit in schizophrenia because this area of research has been
more active than it has been in depression. This is likely because antipsychotic medications
that successfully ameliorate the positive symptoms of schizophrenia (delusions,
hallucinations, etc.) have been available for some time, leaving patients with the residual
negative symptoms, of which a motivation is the primary driver of poor outcome and low
quality of life (Kiang et al. 2003).
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Current concepts of motivation deficits and how motivation is assessed in patients with
schizophrenia is reviewed by Reddy et al. Waltz and Gold extend these concepts into the
exploration of the relationship between a motivation and the representation of expected
value. The clinical research reviewed in the chapters that deal with apathy and motivation in
humans is complimented by a chapter on methods for dissecting motivation and related
psychological processes in rodents (Ward). Research using animal models is critical for
several obvious reasons, including the availability of genetic manipulations, molecular
modifications as well as invasive in vivo monitoring procedures that are not possible in
human subjects. What hasn’t previously been obvious is how well we can use such animal
models to investigate the various components of motivation that are particularly relevant to
human disease. Ward describes such procedures and explains how best to leverage our
current clinical knowledge using state-of-the-art mouse models.
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On the flip side of apathy may be when motivation for a specific goal can come to dominate
action in maladaptive ways as appears to be the case in addictions. Excessive behavior for
many types of rewards including drugs, food, gambling, and sex can be problematic. In
addiction, rapid and strong learning about what leads to reward, excesses in experiencing the
hedonic value of rewards, exaggeration in representing those values, and dominance in being
guided by those representations can all lead to significant narrowing in the diversity of
motives. Several theories exist that attempt to explain the process of addiction in terms of
disruption of motivational processes. Each theory differs in the emphasis on which specific
aspects of motivation are primarily affected. The chapters by both Meyer et al. and Robinson
et al. describe the motivational processes underlying substance abuse disorder. The chapter
by Barrus et al. extends this discussion into the field of gambling. Barrus et al. suggest that
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many of the processes affected in gambling are the same as those affected in drug addiction,
and therefore, the paradigms that have been successfully used to study drug addiction in
animal models can be successfully modified to identify neurobiological mechanisms related
to gambling. The central hypothesis in these analyses of addictions of drugs and gambling
(as well as addiction to food and other things) is that an aberration in reward processing
and/or in the control by cues associated with these rewards underlies the problematic nature
of addictive behavior and its resistance to change.

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8 Treatments
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Given the modern emphasis on reward processes as a fundamental component of motivation,

it is encouraging that modern cognitive/behavioral approaches to treating motivational
disturbances focus on creating reward contingencies that modify deficits or excesses in
behavior. Saperstein and Medalia describe how in schizophrenia patients motivation
enhancing techniques are critical to treatment-related improvements within cognitive
remediation therapy. In the case of addictions, Walter and Petry provide an overview of
research indicating that contingency management is a demonstrably effective psychosocial
treatment for substance use disorders. The central concept of contingency management is
that extrinsic motivators are used to change patients’ behaviors. Specifically, reinforcement
is provided when patients demonstrate abstinence. In the descriptions of both treatment
approaches, the chapters consider the important role that intrinsic motivation may play in
clinical success.
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We are hopeful that the great progress in understanding the neurobiology of motivation
described in this book will influence new ideas that will lead to novel pharmacological,
physiological and psychological/psychosocial approaches to treatments for disorders of
motivation. The identification of novel pharmacological treatments is dependent on the
ability of preclinical researchers to investigate potential targets and screen potential
candidate compounds using truly meaningful endophenotypic assays. The chapters in this
volume that describe clinical studies of patient with disorders of motivation describe how
motivation has been dissected into a number of component processes and the specific
processes that are selectively disrupted in disease have been identified (Reddy et al., Barch
et al., Waltz and Gold). To identify drugs that will be effective for disorders of motivation,
preclinical assays must focus on the same specific processes affected in humans (see Ward in
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this volume). A recent example of the development of the kind of research tools that are
needed for the purpose of investigating potential treatment targets is the strategy of
dissecting goal-directed action from arousal by modifying previously existing rodent
behavioral tasks (Bailey et al. 2015a). These new tools can then be used to assay specific
effects of drugs that affect novel treatment targets (Simpson et al. 2011) and Bailey et al.
(2015b). This novel approach was directly inspired by the literature on the selectivity of
processes disrupted in humans with disorders of motivation.

In addition to pharmacological treatments, there is also the possibility that

electrophysiological treatments for disorders of motivation may be developed. Deep brain
stimulation (DBS) is currently used to treat a number of neurological and psychiatric
conditions (Kocabicak et al. 2015; Kringelbach et al. 2007; Udupa and Chen 2015). DBS
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has been used to treat essential tremor, Parkinson’s disease, treatment refractory major
depression, severe obsessive–compulsive disorder, and chronic pain for several years.
Several other applications are in experimental stages, including clinical trials for pervasive
addiction and symptoms of schizophrenia. Such an invasive procedure requires many
successive small-scale clinical trials before optimal procedures can be successfully
developed. A significantly less invasive procedure used to modulate brain activity is
transcranial magnetic stimulation (TMS). While the mechanism(s) by which TMS alters
neuronal function and network activity is not understood, due to its noninvasiveness,

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hundreds of clinical trials have been conducted for a long list of neuropsychiatric conditions,
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including schizophrenia, and craving/addiction. A comprehensive review of studies of

repetitive TMS has recently been published (Lefaucheur et al. 2014).

Lastly, we are also hopeful that the emerging understanding that there are multiple systems
driving motivation on an organismal level will lead to the development of treatment schemes
that are more comprehensive than those that have been developed in the past. It may be that
subtle adjustments in several of the factors that are involved in disorders of motivation (the
endocrine system, circadian system, neurotransmitter function, etc.) can result in greater
improvements and less side effects than treatments that focus on a single system.

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Fig. 1.
A simplified diagram of the influencing factors and processes that are involved in
motivation. This framework of motivation places cost–benefit analysis central to the concept
of motivation. Three major categories of factors are known to influence motivation: the
individual’s physiological state, the environment, and the individual’s past history.
Information about all 3 categories of factors will be subject to a number of processes
(represented inside the blue oval), including evaluation and encoding. In almost all
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circumstances, the motive, environment, and physiological state will not be novel; therefore,
information will also undergo learning and retrieval processes. All of the combined
processes result in weighting of all the costs and benefits related to the motive, and the
output of the cost–benefit calculation will impact upon the direction and vigor of action that
the individual takes toward the motive goal
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